Mid dle and Late Pleis to cene ter res trial snails from the Mid dle Dniester area, Ukraine (based on Mykola Kunytsia’s col lec tions)
Yana POPIUK1, Bogdan RIDUSH1, * and Tatiana SOLOVEY2
1 Yuriy Fed’kovych Chernivtsi Na tional Uni ver sity, Kotsubynskogo 2, 58012 Chernivtsi, Ukraine
2 Pol ish Geo log i cal In sti tute – Na tional Re search In sti tute, Rakowiecka 4, 00-975 Warszawa, Po land
Popiuk, Y., Ridush, B., Solovey, T., 2021. Mid dle and Late Pleis to cene ter res trial snails from the Mid dle Dniester area, Ukraine (based on Mykola Kunytsia’s col lec tions). Geo log i cal Quar terly, 65: 6, doi: 10.7306/gq.1575
Ex ten sive col lec tions of land molluscs from the many sec tions of Mid dle and Late Pleis to cene de pos its in the re gion of the Mid dle Dniester River were made by Pro fes sor Mykola Kunytsia (1925–2002). These col lec tions, now at the Nat u ral Mu - seum, Yuriy Fedkovych Chernivtsi Na tional Uni ver sity, were for the most part ana lysed, but re mained par tially un pub lished.
M. Kunytsia used them for re gional re con struc tions of palaeolandscapes dur ing var i ous stages of the Pleis to cene. In our study, we used these col lec tions to as sess the land snail di ver sity in the re gion dur ing con sec u tive stages of the Mid dle and Late Pleis to cene. Our anal y sis of fau nas of cold and warm stages showed that the lat ter were more di verse. While all or most of the cold stages had many spe cies in com mon, the fau nas of the warm stages were more het er o ge neous. Such a fea ture may play an es sen tial role in Qua ter nary biostratigraphy.
Key words: Mid dle Dniester, ter res trial molluscs, biodiversity, Pleis to cene.
INTRODUCTION
Due to its nu mer ous well-ex plored Qua ter nary se quences, the Mid dle Dniester re gion can be re garded as on one of the key ter ri to ries for Qua ter nary stud ies in East ern Eu rope. The re gion cov ers the can yon-like part of the Dniester River val ley, ap prox - i mately be tween towns of Halych and Rybnytsia, in clud ing can - yon-like sec tions of the Dniester trib u tar ies. In this re gion, the Dniester val ley is quite deep (270–300 m), and wide, with a well-de vel oped suc ces sion of an cient river-ter races. Most of the Late and Mid dle Pleis to cene ter races are lo cated in side the deep and nar row can yon-like part of the val leys; how ever, some high ter races of the Early Pleis to cene and Plio cene are wide and de vel oped out side the can yon (Ridush and Marchuk, 2018).
Dif fer ent au thors have iden ti fied var i ous num bers of ter - races in the Dniester val ley, from 6 to 13 (Tomeniuk, 2010).
Based on a palaeopedological method, Veklitch (1982) dis tin - guished 16 ter races above the mod ern floodplain. His ap proach was adopted by the State Geo log i cal Sur vey of Ukraine.
The gen eral pat tern of the Dniester val ley looks like a se ries of huge, deeply in cised me an ders. As a rule, the in ner-can yon Pleis to cene ter races, num bered from 1st to 10th, and from 10 to 100 m high above the mod ern wa ter ta ble, are pre served on
the con vex sides of the me an ders, while the con cave bank of the river is usu ally steep. Over all, these ter races have a two-fold struc ture, con sist ing of an al lu vial suite, rest ing on the bed rock base ment, and a sub-ae rial suite, com pris ing slope and ae olian de pos its. Both al lu vial and sub-ae rial suites con tain rich fau nas of ter res trial snails.
The mid dle sec tion of the Dniester River val ley is fa mous for its nu mer ous Palaeo lithic sites as so ci ated with the Pleis to cene loess-palaeosol se quences of the an cient river ter races (e.g., Goretsky and Tzeitlin, 1977; Goretsky and Ivanova, 1982;
Ivanova and Tzeitlin, 1987; Anisyutkin, 2013; Kulakovska et al., 2015; £anczont and Madeyska, 2015). Al most all strata of these se quences con tain shells of ter res trial molluscs which have tra di tion ally been used for the palaeo eco logi cal re con - struc tions. A num ber of schol ars have stud ied the Pleis to cene snail fau nas since 1880 (Dunikowski, 1880; B¹kowski, 1880, 1881, 1884, 1885, 1891; £omnicki, 1886, 1887, 1900, 1908;
Teisseyre, 1900; Friedberg, 1906; Wiœniewski, 1908; Rogala, 1907; Rychlicki, 1913; Polianskyi, 1925; Petrbok, 1930;
Ambrozewicz, 1932, 1938; Lungersgauzen, 1933, 1938;
Bondarchuk, 1933, 1959; Danilovskyi, 1940, 1961; Berg, 1946;
Radzijevskyi, 1957, 1959; Veklitch, 1961, 1968; Ivanova, and Popov, 1961; Kunytsia, 1964, 1965, 1966, 1968, 1969, 1971, 1974, 1975, 1978; Melnychuk, 1972, 1984, 2004; Motuz, 1977, 1982, 1987; Dmytruk, 1998, 2000, 2001).
The great est con tri bu tion to the study of the Pleis to cene malacofauna of the re gion was made by prof. Mykola Kunytsia (1925–2002). His sum ma ris ing re search, which cov ered al most all the ter ri tory of Ukraine, was pub lished a few years af ter his death. He col lected and stud ied molluscs of the dif fer ent Pleis - to cene lithological-strati graphic units, iden ti fied the spe cies, pro vided a palaeontological-strati graphic cor re la tion and traced
* Cor re spond ing au thor, e-mail: b.ridush@chnu.edu.ua Re ceived: July 6, 2020; ac cepted: No vem ber 8, 2020; first pub lished on line: January 25, 2021
zonal-geo graph ical changes of the malacofauna. He was the first to re con struct palaeolandscapes and palaeoclimates of the gla cial and inter gla cial phases based on eco log i cal-zoo geo - graph i cal anal y sis and to pre pare palaeolandscape maps of Ukraine (Kunytsia, 2007). Most of Kunytsia’s sites were lo cated in the Mid dle Dniester re gion.
Most of pre vi ous stud ies of the mol lusc as sem blages of the re gion had used them for palaeo eco logi cal re con struc tions, as tem per a ture and hu mid ity in di ca tors. Our pa per fo cuses on their spe cies di ver sity, and on iden ti fy ing com mon or dis tinc tive fea tures of the Mid dle and Late Pleis to cene mol lusc fau nas of the re gion.
MATERIAL AND METHODS
The mol lusc di ver sity anal y sis was based on the abun dant ma te rial pub lished by Kunytsia (1965, 1966, 1971, 1974, 1984,
and 2007), as well as on his un pub lished col lec tions. The list of spe cies was sup ple mented with data of other re search ers (Motuz, 1977, 1982, 1987; Melnychuk, 1984, 2004; Dmytruk, 2004; Bogucki et al., 2011, 2012; Alexandrowicz et al., 2014;
Alexandrowicz, 2015a, b).
In the biodiversity anal y sis, in the de ter mi na tion of changes in spe cies com po si tion at dif fer ent stages of the Pleis to cene (Ta ble 1), and in the in ter pre ta tion of palaeo eco logi cal con di - tions, we spec i fied the most com mon spe cies which were pres - ent at more than one site. To avoid in ter pre ta tion er rors and to ex clude ran dom finds, we ex cluded from the anal y sis spe cies which were rep re sented at sin gle sites only. Such finds may in - di cate lo cal dif fer ences or re-de po si tion and re quire fur ther ver i - fi ca tion.
De spite the great num ber of pub li ca tions deal ing with the malacofauna of the re gion, not all of them spec ify the ex act strati graphic po si tion of the sam ples, and this is of ten true also of Kunytsia’s re search. In some other pub li ca tions, some spe -
T a b l e 1 Main strati graphic units of the Mid dle and Late Qua ter nary in East ern and West ern Eu rope (com piled af ter Lindner et al., 2004;
Lisiecki and Raymo, 2005; Gerasimenko, 2010a; Velichko et al., 2011)
* The age in ka is given for stages af ter Gerasimenko (2010a) and af ter Lisiecki and Raymo (2005) for MIS
cies are largely as signed to MIS 2-4 (Alexandrowicz et al., 2014). We used that part of the ma te rial which was un am big u - ously as signed to par tic u lar strati graphic lev els and ages within the de pos its.
The com par a tive anal y sis and as sess ment of biodiversity of the Pleis to cene mol lusc as sem blages was car ried out sep a - rately for cold and warm stages. It was found that the fre quent cli mate changes led to eco log i cal and geo graph ical vari abil ity of snail as sem blages at dif fer ent stages. Pri mar ily, this con cerns changes in the spe cies com po si tion of mol lusc as sem blages in the warm and cold stages of the Pleis to cene (glacials and interglacials).
The fau nas of the cold and warm stages dif fered sig nif i - cantly. How ever, the spe cies’ com po si tion could also vary among the fau nas of warm and cold phases. Such vari a tions de pended mainly on the tem per a ture, hu mid ity, du ra tion of in di - vid ual cold or warm stages, and the cy clic na ture of cli mate changes within those stages, as well as on the abil ity of some spe cies to sur vive cli mate change, adapt, and re ap pear in sub - se quent stages as rel ict forms. The in ter ac tion of all these fac - tors should be re flected in the in di vid ual fea tures of mol lusc as - sem blages at dif fer ent stages. Be cause of this, we con sid ered it ap pro pri ate to com pare the di ver sity of molluscs sep a rately in cold and warm stages. It is such com par i sons that should al low us to spec ify the dif fer ences in the tax o nomic and eco log i cal com po si tion of small-scale units of dif fer ent ages.
Eco log i cal clas si fi ca tion of the spe cies was based on Alexandrowicz and Alexandrowicz (2011), de tail ing malacological anal y sis, re search meth ods, pro cess ing, and in - ter pre ta tion of subfossil mol lusc as sem blages. The in ferred en - vi ron men tal con di tions of the Pleis to cene stages based on the mol lusc as sem blages of the re gion were com pared with the re - sults of palaeo-palynological re con struc tions of the Pleis to cene (Gerasimenko, 2010a).
Apart from Kunytsia’s pub li ca tions, much ma te rial ac cu mu - lated dur ing his life time long re mained un pub lished. Most was de pos ited at the Nat u ral Mu seum, Yuriy Fedkovych Chernivtsi Na tional Uni ver sity, more than 15 years ago. They are of con - sid er able value and ana lys ing them is im por tant for Qua ter nary malacology. Some of the ma te rial orig i nates from land now flooded by the Dniester res er voir, and so their pres ence and sig nif i cance are es pe cially rel e vant. We are the first to ana lyse Kunytsia’s un pub lished ma te ri als. The work was car ried out in sev eral stages, which in cluded cat a logu ing of the ma te rial, its curation and iden ti fi ca tion. Over all, the col lec tion con sists of 897 glass tubes con tain ing mol lusc shells from dif fer ent re gions of Ukraine, which we sorted by lo cal ity. The most abun dant ma - te rial come from the Mid dle Dniester val ley, and only molluscs from that area are de scribed in this pa per.
The la bels mostly per tain to the geo graph ical lo ca tion (near - est town or vil lage in whose vi cin ity the sam pled geo log i cal sec - tion was sit u ated); sed i ments in the sec tions; mol lusc spe cies iden ti fied; rarely, de tailed geo graphic ref er ence of the sec tion rel a tive to the town or vil lage; and depth of sam pling in the sec - tion. Over all, more than 458 glass tubes with subfossil ma te rial are from the mid dle part of the Dniester val ley. Un for tu nately, many of the tubes do not bear any in di ca tion of the lo ca tion, or just give the num ber of the sec tion (for ex am ple, sec tion 226 –
‘P. 226’). The in for ma tion re gard ing such sam ples was con - tained in Kunytsia’s di a ries, which are lost.
We were able to re con struct in for ma tion con cern ing a num - ber of lo cal i ties (Fig. 1; 1–43); these in clude sec tions in the vil - lages of Ataky (19 sam ples), Rukhotyn (5 sam ples), Gorodok (6 sam ples), Dnistrove (8 sam ples), Isakivtsi (5 sam ples), Zavallya (2 sam ples), Kudryntsi (4 sam ples), Kamyanka (4 sam ples), Naddnistrianka (26 sam ples), Smotrych (27 sam -
ples), Teremtsi (8 sam ples), Stara Ushytsia (15 sam ples), Molodove (29 sam ples), Repuzhyntsi (29 sam ples), Zalishchyky (30 sam ples), Ozhevo (59 sam ples), Lypchany (54 sam ples), Vasylivka (20 sam ples), Voloshkovo (45 sam - ples), Nezvysko (19 sam ples) and Halych (31 sam ples).
The fol low ing spe cies are rep re sented in the sam ples from these lo cal i ties: Acicu la polita, Carychium min i mum, Succinea oblonga, S. o. elongata, S. putris, Cochlicopa lubrica, C. nitens, Acanthinula aculeata, Vallonia costata, V. pulchella, V. enniensis, V. tenuilabris, Pupilla muscorum, P. loessica, P. sterri, Ver tigo pygmaea, V. parcedentata, V. angustior, Columella edentula, C. columella, Orcula dolium, Chondrula tridens, Cochlodina laminata, Clausilia dubia, Cl. cruciata, Laciniaria plicata, Vitrea crystallina, Nesovitrea hammonis, Euconulus fulvus, Trichia hispida, Helicopsis striata, Xerolenta obvia (= Helicella candicans), Perforatella bidentata, Pseudotrichia rubiginosa, Euomphalia strigella, Cepaea vindobonensis, Helix ðomatia, Theodoxus fluviatilis, Fagotia acicularis, Bithynia tentaculata, Lithoglyphus naticoides, Lymnaea (Stagnicola) palustris, Physa fontinalis, Anisus spirorbis, Pisidium casertanum. Many sam ples have not yet been iden ti fied and re quire fur ther re search.
Since the strati graphic po si tion of some sam ples re mains un known, not all spe cies from the col lec tion are in cluded in our anal y sis. How ever, in for ma tion on the lo ca tion of the sec tions makes it pos si ble to re-sam ple the sites in the fu ture, to re fine the data.
RESULTS
The Pleis to cene cli mate changes in volved a suc ces sion of cold and warm phases of dif fer ent am pli tude. It is ob vi ous that the mol lusc fau nas of cold and warm phases were quite dif fer - ent, but even mol lusc as sem blages among the cold and warm phases var ied con sid er ably; that is why we ana lyse the mol lusc di ver sity sep a rately for the cold and warm phases.
An es sen tial fea ture of molluscs dur ing the cold phases is their ad ap ta tion to cli mate changes, as ex pressed by mor pho - log i cal changes in the shell size and struc ture: ap er ture size, shell thick ness, pres ence of ap er tural bar ri ers etc. As noted by Kunytsia (1966), the shells of ubiq ui tous spe cies of pro longed cold phases dif fer from those of warm stages in marked traces of ad verse hab i tat ef fects. For ex am ple, ter res trial spe cies have smaller shells (some be ing dwarfed) com pared with typ i cal spec i mens, pos sess a di min u tive ap er ture, are thin-walled in ar eas close to the ice front and thick ened on cal cium-rich cal - car e ous de pos its. Fresh wa ter snails in cold con di tions are rel a - tively small and have thick ened shell walls. Such vari a tion serves as an ef fec tive ba sis for the re con struc tion of cli mate con di tions not only in the case of stenoecious spe cies, but also for euryoecious snails. Cli mate changes were also re flected in the spe cies com po si tion of the Pleis to cene mol lusc as sem - blages and their geo graph ical dis tri bu tion at dif fer ent cli ma tic stages. All these pat terns are well-re flected in the as sem blages from the Mid dle Dniester re gion.
Within this re gion, spe cies as sem blages of dif fer ent warm phases are more dif fer ent than those of dif fer ent cold phases, while in di vid ual fea tures are clearly man i fested in some in ter - vals. This spec i fic ity of inter gla cial fau nas has also been noted for north west ern Eu rope (Limondin-Lozouet and Preece, 2014). This is due to the more sig nif i cant vari abil ity of the land - scapes dur ing the warm stages.
Our stud ies show that the ubiq ui tous Succinea oblonga was com mon through out the Pleis to cene, both in the cold/warm, and in the dry/wet phases. Typ i cal cold-lov ing spe cies which
oc curred through out all the cold phases of the Pleis to cene are Columella columella and Vallonia tenuilabris (Kunytsia, 1965, 1966, 1971, 1974, 1984, 2007; Motuz, 1977, 1982, 1987;
Melnychuk, 1984, 2004; Dmytruk, 2001; Bogucki et al., 2002, 2004; Alexandrowicz et al., 2014). Xerolenta obvia (= Helicella candicans) is thermophilous. How ever, it some times ap pears in the cold in ter vals, and this is clearly due to the arid ity be cause it is also as so ci ated with xe ric hab i tats. De spite the com par a tively uni form com po si tion of snail as sem blages of the cold phases, there are some phase-to-phase dif fer ences.
Malacofaunas of the cold stages of the Mid dle and Late Pleis to cene and their di ver sity
The Sula Stage (650–600 ka, MIS 18) is dis tin guished by a con sid er able spe cies di ver sity (Ta ble 2). Typ i cal spe cies of this stage do not oc cur in the suc ceed ing cold Pleis to cene pe ri ods:
Ver tigo aff. ronnebyensis, Vallonia pulchella, V. enniensis, Cochlicopa lubrica, Nesovitrea pet ro nella. Two spe cies in the mid dle Dniester val ley, Ver tigo alpestris and Ver tigo genesii, have been found only in Sula and Bug loesses.
The fauna of the Sula stage has a spe cific eco log i cal and zoo geo graph i cal com po si tion. The as sem blage in cludes spe - cies of dif fer ent hab i tat pref er ences, while mostly cold-lov ing or cold-re sis tant. Be sides the pre dom i nance of meso philes, hygrophiles and psy chro philes are pres ent (Ver tigo genesii, Vallonia enniensis, Pseudotrichia rubiginosa); they pre fer moist ar eas and pos si bly even wetlands. Alexandrowicz (2015a) noted that as sem blages with Ver tigo genesii are char ac ter ized by the pres ence of many moist-lov ing and cold-lov ing spe cies.
The fauna of this stage in cludes also nu mer ous xerophiles (Pupilla sterri, P. triplicata, Chondrula tridens, Xerolenta obvia Fig . 1. Pleis to cene sites with mol lusc fau nas in the Mid dle Dniester val ley
(ac cord ing to un pub lished ma te ri als by M. Kunytsia)
1 – Halych, 2 – Koropets, 3 – Nezvysko, 4 – Zalishchyky, 5 – Repuzhyntzi, 6 – Dobrovliany, 7 – Horodok, 8 – Onut, 9 – Rukhotyn, 10 – Dnistrove, 11 – Kudryntsi, 12 – Zavallya, 13 – Isakivtsi, 14 – Ataky, 15 – Hotyn, 16 – Smotrych, 17 – Kamianka, 18 – Voronovytsia, 19 – Makarivka, 20 – Nagoriany, 21 – Hrushivtsi, 22 – Lenkivtsi, 23 – Babyn, 24 – Dnistrivka, 25 – Molodove, 26 – Korman, 27 – Stara Ushytsia, 28 – Neporotove, 29 – Lomachyntsi, 30 – Naddnistrianske, 31 – Zhvan, 32 – Bernashivka, 33 – Lypchany, 34 – Ozheve, 35 – Vasylivka, 36 – Rozpopyntsi, 37 – Voloshkovo, 38 – Yaryshiv, 39 – Mogyliv-Podilskyi, 40 – Bila, 41 – Porogy, 42 – Kosoutsy, 43 – Velyka Kisnytsia
(= Helicella candicans); there are many spe cies of open hab i - tats (Ver tigo parcedentata, Pupilla muscorum, P. densegyrata, P. sterri, P. triplicata, Vallonia pulchella, V. tenuilabris), and typ i - cal for est-dwell ers (Ver tigo aff. ronnebyensis). Such a di verse set of spe cies in the con text of eco log i cal con di tions sug gests that dur ing the 50 ky of the Sula Stage (650–600 ka) (Gerasimenko, 2010a), within the Mid dle Dniester area, sev eral as sem blages suc ces sively de vel oped, of ten re garded as one taphocoenosis. The ver ti cal fa cies vari a tion of the de pos its, and of those as sem blages, shows that the be gin ning of the Sula Stage was rather cold and rel a tively hu mid, and its later phase was cold and dry (Kunytsia, 1974). This is sup ported by the palynological data, ac cord ing to which two cold sub-stages (sl1
and sl2) and one rel a tively warm stage (sl3) oc curred in Sula time (Gerasimenko, 2010a).
The Tyligul fauna (500–410 ka, MIS 12) is typ i cal of Pleis - to cene loess de pos its (Ta ble 2). An ex cep tion is the pres ence of Oxyloma (Succinea) elegans, which re flects par tic u lar palaeo-geomorphological con di tions, of ten be ing found on floodplains. Pupilla sterri, which is pres ent at all other cold stages of the Mid dle and Late Pleis to cene, was not found in the Tyligul set. In gen eral, the fauna is rep re sented mainly by open-hab i tat spe cies (Ver tigo parcedentata, Pupilla muscorum, P. triplicata, Vallonia tenuilabris). Its bulk is com posed of meso- and xerophiles. There is a large group of cryophilic spe cies (Columella columella, Vallonia tenuilabris, Ver tigo parce den - tata). This as sem blage was char ac ter is tic of the Pleis to cene cold steppe (tun dra-steppe; Kunytsia, 2007). De spite the ab - sence of pro nouncedly shade-lov ing spe cies, it in cluded such spe cies as Euconulus fulvus and Pseudotrichia rubiginosa which pre fer for ested or par tially shaded wet hab i tats. Some - times they in habit dry hab i tats, but still with mod er ately rich veg - e ta tion (Moine et al., 2005). Pseudotrichia rubiginosa is also clas si fied as a psy chro phile, in hab it ing mostly wet or moist for - ests.
Mainly cold-lov ing and cold-re sis tant spe cies of open dry hab i tats (Ta ble 2) are typ i cal of the Dnie per Stage (180–127 ka, MIS 6, Saalian): Ver tigo parcedentata, Columella columella, Pupilla muscorum, P. sterri, P. triplicata, P. loessica, Vallonia tenuilabris, Xerolenta obvia (= Helicella candicans), Helicopsis striata. Psychro- and hygrophiles are rep re sented by Ver tigo genesii (Alexandrowicz et al., 2014) and Zonitoides nitidus (Melnychuk, 2004).
Pseudotrichia rubiginosa, which was pres ent at all other cold stages of the Pleis to cene, is ab sent from the Dnie per as - sem blages. How ever, here for the first time ap pears Helicopsis striata, which is pres ent in all sub se quent cold stages. It is not known from ear lier cold stages, but in the Mid dle Dniester the spe cies also ap pears in the Mid dle Pleis to cene as sem blages of the warm Zavadivka time.
Ac cord ing to Melnychuk (2004), Pupilla loessica and Zonitoides nitidus have been found in the Dnie per de pos its, and are known also from the Bug loesses. Ac cord ing to Kunytsia (2007) Pupilla loessica is an in dex form for the last gla ci ation. Its rare finds in the de pos its of the Dnie per Stage are known from the Dnie per val ley (Vasyl’kiv). The spe cies is typ i cal of dry cold Pleis to cene stages. It is re garded as an in di ca tor of ac tive ae - olian loess ac cu mu la tion (Ložek, 1964). For a long time, it was con sid ered ex tinct (Kunytsia, 2007), but now it is known from the re cent fauna in the moun tain ous re gions of Cen tral Asia (Ložek, 1986; Meng and Hoffmann, 2009).
Mol lusc as sem blages of the Tyasmyn Stage (110–104 ka) are rel a tively spe cies-poor (Ta ble 2). The most char ac ter is tic
cold-lov ing meso philes are Columella columella and Vallonia tenuilabris; xerophiles of open hab i tats in clude Pupilla sterri, Helicopsis striata, and Xerolenta obvia (= Helicella candicans).
The psy chro phile Pseudotrichia rubiginosa is less com mon.
The de pos its do not con tain the cryophile Ver tigo parcedentata, which is pres ent in as sem blages of all the other cold stages of the Pleis to cene. The xerophile Pupilla triplicata, which was found in the de pos its of all the pre vi ous cold stages, also dis ap - peared. Later it ap pears only at the end of the Pleis to cene, in the Prychornomorya Stage (see be low).
The fauna of the Uday Stage (74–55 ka, MIS 4) is mainly com posed of xero- and meso philes of open hab i tats, which are of ten found in other cold pe ri ods of the Pleis to cene (Ta ble 2).
The char ac ter is tic cryophylic spe cies are pres ent. Vitrea crystallina, which in hab ited the area at that time, is spe cific for this stage. This is a typ i cal rep re sen ta tive of shade-lov ing group (Alexandrowicz and Alexandrowicz, 2011) and was not de - tected in the other cold stages. It pre fers cool and wet con di - tions (Moine et al., 2005). The xerophile Helicopsis instabilis ap pears here for the first time; it also oc curs in the fol low ing cold stages of the Pleis to cene.
Among all other stages of the Pleis to cene (cold and warm), the mol lusc as sem blages of the cold est Bug Stage (27–18 ka, MIS 2) show the great est tax o nomic and eco log i cal di ver sity (Ta ble 2). Be sides the more wide spread spe cies, at that time the mid dle part of the Dniester val ley was in hab ited by: the cryophiles Columella columella, Vallonia tenuilabris, Ver tigo parcedentata; typ i cal spe cies of open hab i tats Pupilla muscorum, P. loessica, Vallonia excentrica, etc.; the meso - philes Cochlicopa lubrica, Ver tigo alpestris, V. arctica, V. pseudosubstriata, Trichia hispida; xerophiles Pupilla sterri, P. triplicata, Chondrula tridens, Xerolenta obvia (= Helicella candicans), Helicopsis striata, Helicopsis instabilis; and the psychro- and hygrophiles Ver tigo genesii, Zonitoides nitidus, Pseudotrichia rubiginosa.
Vallonia excentrica, Ver tigo arctica and V. pseudo - substriata dur ing the Pleis to cene are found only in Bug time; to - day they in habit tun dra and marshy hab i tats of Scan di na via, the Brit ish Isles and Ice land. Ver tigo arctica is also dis trib uted in the Tatra Moun tains and the Aus trian Alps (Nekola et al., 2018).
Ver tigo alpestris has been found in the cold phases of the Bug and Sula time only; V. genesii was found in the de pos its of the Bug, Dnie per, and Sula times. Shells of Pupilla muscorum densegyrata in the Mid dle Dniester re gion are known only from the Bug Stage, though in gen eral Pupilla muscorum is also char ac ter is tic of many cold stages.
Such a great taxomomic, quan ti ta tive, eco log i cal, and geo - graph ical rich ness of the fauna dur ing a rel a tively short time (com pared to other cold stages), which was the cold est time in the Pleis to cene, de serves spe cial at ten tion. Ob vi ously, in cer - tain palynologicaly es tab lished sub-stages (cryo-hygrotic bgd+e
and cryo-xerotic bgf; Gerasimenko, 2010a), when the palaeolandscapes changed sig nif i cantly, some mol lusc as sem - blages were re placed by oth ers. Such a di ver sity of molluscs re - flects the land scape-cli mate changes of the Bug Stage.
Some rel ict cryophiles sur vived to the Prychornomorya Stage (15–10 ka, MIS 2; Ta ble 2). The pro por tion of hydrophiles and mesophilous for est-dwell ers in creased.
Clausilia dubia, Cl. bidentata, Arianta arbustorum, which are as so ci ated with coldstorms (Moine, 2005), ap pear for the first time in the cold stages. Nesovitrea hammonis, which was known from Sula time, re ap pears.
T a b l e 2 Changes in the spe cies di ver sity of ma jor molluscs dur ing the cold stages of the Pleis to cene in the Mid dle Dniester area
(af ter Kunytsia, 1965, 1966, 1971, 1974, 1984, 2007; Motuz 1977, 1982, 1987; Melnychuk, 1984, 2004; Dmytruk, 2004;
Ivchenko, 2010; Bogucki et al., 2011, 2012; Alexandrowicz et al., 2014)
MALACOFAUNAS OF THE WARM STAGES OF THE MIDDLE AND LATE PLEISTOCENE
AND THEIR DIVERSITY
Com pared with the cold stages, the mol lusc as sem blages of warm stages dif fer more among them selves. The as sem - blages of the last warm stages, Pryluky, Vytachiv, and Dofinivka (Ta ble 1), are richer in spe cies. The lat ter is dis tinct in hav ing relic cryophilic spe cies.
The as sem blages of the Martonosha Stage (780–650 ka, MIS 19-17) are spe cies-poor (Ta ble 3). Be sides the ubiq ui tous forms, Vallonia pulchella and Chondrula tridens, rep re sent ing open dry hab i tats and wide spread in the warmer stages, were found.
In hab it ants of open dry hab i tats (Pupilla muscorum, Vallonia pulchella, V. costata, Chondrula tridens, Xerolenta obvia (= Helicella candicans) are char ac ter is tic of the Lubny Stage (600–500 ka, MIS 15-13). The meso- and psy chro philes are rep re sented only by Carychium min i mum and Punctum pygmaeum; this sug gests that dur ing this pe riod, the area was in hab ited by spe cies of warm steppes and only the last two spe - cies pro vide ev i dence of pe ri ods of higher hu mid ity (Ta ble 3).
Ac cord ing to Gerasimenko (2010a), the Lubny Stage con - sists of three sub-stages: two of these were mod er ately warm, with a sig nif i cant south ward ex pan sion of for ests (some times up to the steppe bound ary), and a cold in ter val sep a rat ing them.
How ever, by the end of each warm sub-stage, for ests were re - placed by meadow-steppe land scapes. The eco log i cal com po - si tion of the as sem blages which in hab ited the Mid dle Dniester val ley at that time in di cates lon ger dry phases, as shown by the pres ence of nu mer ous dry-lov ing spe cies. At the same time, it can not be ruled out that the de pos its of wet ter stages with their fauna were sim ply not pre served, or have not been found yet, since cold-lov ing spe cies in the fauna are also ab sent, and in sub-stage lb2, along with the for ma tion of thin loess/loam in sub-periglacial land scapes, two gen er a tions of pri mary ground
vines had also de vel oped (Gerasimenko, 2010a). There fore, the ex ist ing mol lusc ma te ri als of the Lubny Stage should be re - vised.
In Zavadivka time (410–250 ka, MIS 11-7) the area was in - hab ited mainly by open-ground spe cies (Euomphalia strigella), as well as xerophiles: Pupilla sterri, Helicopsis striata, and He lix lutescens. Aegopinella nitens and He lix pomatia rep re sent spe - cies of for est or well-shaded hab i tats. The mesophile Trichia hispida, wide spread in the warm stages of the Pleis to cene, was also pres ent. Vallonia pulchella and Chondrula tridens, char ac - ter is tic of the other warm stages, are ab sent from the de pos its of the Zavadivka and Dofinivka stages. These spe cies pre fer open and dry hab i tats; other mem bers of this eco log i cal group are char ac ter is tic. The spe cies com po si tion of the these stages is sig nif i cantly dif fer ent since in the Dofinivka there is a sig nif i - cant pro por tion of cryophiles, which are not found in the Zavadivka (Ta ble 3).
The de pos its of the Kaydaky Stage (130–110 ka, MIS 5e) con tain mainly spe cies which are typ i cal of most of the warm stages of open dry hab i tats: Vallonia pulchella, Chondrula tridens, Xerolenta obvia (= Helicella candicans). Vallonia costata rep re sents the same eco log i cal group, but it is less com mon in the warm stages (only in the Lubny and Kaydaky).
The hygrophile Succinea (Oxyloma) elegans, which lives in wet hab i tats close to wa ter, was found only in the de pos its of the Kaydaky Stage. Its pres ence sug gests floodplain con di tions dur ing de po si tion at those lo cal i ties (Ta ble 3).
The as sem blage of the Pryluky Stage (104–74 ka, MIS 5a-c) is the rich est in spe cies among the warm stages (Ta - ble 3). It is also di verse in its eco log i cal com po si tion. The typ i cal rep re sen ta tives of the light-lov ing and xerophilous fauna are Pupilla sterrii, Vallonia pulchella, Chondrula tridens, Xerolenta obvia (= Helicella candicans), Helicopsis striata. The group of shade-lov ing forms is also nu mer ous: Clausilia dubia, Vitrea diaphana, Arianta arbustorum, Cepaea aff. nemoralis, He lix pomatia. The as so ci a tion of this warm stage also in cludes the Tabl. 2 cont.
T a b l e 3 Changes in the spe cies di ver sity of ma jor molluscs dur ing warm Pleis to cene stages in the Mid dle Dniester area (af ter Kunytsia, 1965, 1966, 1971, 1974, 1984, 2007; Motuz, 1977, 1982, 1987; Melnychuk, 1984, 2004; Dmytruk, 2004;
Ivchenko, 2010; Bogucki et al., 2011, 2012; Alexandrowicz et al., 2014)
Pleis to cene cryophile Vallonia tenuilabris. Trichia hispida and Clausilia parvula rep re sent the mesophilous group. The spec i - fic ity of the as sem blage of the Pryluky Stage is the pres ence of the mesophile Clausilia parvula. In the Mid dle Dniester re gion, it is known only from the next warm Vytachiv Stage. At pres ent, the spe cies in hab its West ern and Cen tral Eu rope, but the dis tri - bu tion range reaches the fringes of the West ern Carpathians. It has been found and stud ied in de tail in Po land, where it be longs among rare mod ern spe cies (Maltz, 1999; Pokryszko et al., 2004; Pokryszko and Cameron, 2005). It is a mesophile, petrophile and calciphile (Pokryszko and Maltz, 2007). It is char ac ter is tic of open, mod er ately moist ar eas, and is of ten found in rocky hab i tats, es pe cially on veg e ta tion-cov ered lime - stone and rarely in for est leaf-lit ter. At pres ent, this spe cies does not live in Ukraine (Likharev, 1962).
The fauna of the Vytachiv Stage (55–27 ka, MIS 3) is also quite rich in spe cies (Ta ble 3). The great est pro por tion is formed by dry-lov ing spe cies of open ar eas: Pupilla muscorum, P. sterri, Vallonia pulchella, Chondrula tridens, Xerolenta obvia (= Helicella candicans), Helicopsis striata. Shade-lov ing spe - cies (Clausilia dubia, Vitrea diaphana, Arianta arbustorum) and meso philes (Clausilia parvula, Trichia hispida) also con sti tute a sig nif i cant pro por tion of the as sem blage. Most of the spe cies listed are typ i cal of the warm Pleis to cene stages.
The de pos its of the short Dofinivka Stage (18–15 ka, MIS 2), hold many cold-lov ing spe cies (Ver tigo parcedentata, Columella columella, Vallonia tenuilabris) which are not char ac - ter is tic of warm pe ri ods. Xerophiles of open ar eas are rep re - sented by Pupilla sterrii, Xerolenta obvia (= Helicella candicans) and Helicopsis striata. The shade-lov ing group forms a con sid - er able pro por tion: Cochlodina laminata, Clausilia dubia, Arianta arbustorum; the same is true of meso- and psy chro philes – Laciniaria plicata, Trichia hispida, Pseudotrichia rubiginosa (Ta - ble 3). The as sem blage is quite di verse for the rather short du - ra tion of the Dofinivka Stage. It in di cates that at that time the cli - mate con di tions and land scapes in the re gion of the Mid dle Dniester of ten changed. In dif fer ent sec tions, de pos its of the stage are rep re sented by steppe-type brown soils, cher - nozem-like soils, and grey for est soils. In the sec tion close to Koropets vil lage (Figs. 1 and 2), two soils can be traced: a lower cher nozem and an up per grey for est soil, which are sep a rated by inter-soil loams con tain ing cold-lov ing fauna (Kunytsia, 1974).
The mol lusc as sem blages and the de pos its of the Dofinivka Stage are a vivid ex am ple of Pleis to cene en vi ron men tal-cli mate changes, that could lead to a dras tic change of land scapes within rather short time in ter vals, and of how the mol lusc fauna, as an ex cel lent eco log i cal in di ca tor, re acted to these changes by re plac ing of some spe cies (as sem blages) with oth ers. A sim -
i lar phe nom e non can be ob served in the Ho lo cene, when dur - ing 12 ky the en vi ron men tal con di tions changed re peat edly.
Ana lys ing the late stages of the Pleis to cene, and the na ture and fre quency of these changes, the ques tion arises: “Could such changes have taken place dur ing the more an cient stages of the Pleis to cene, which lasted 50–100 ky?” To an swer this, it is nec es sary to study in de tail the depositional se quence of such stages, the se quence and changes in their suc ces sion, fa cies vari abil ity and, cru cially, to sam ple molluscs strictly in ac cor - dance with the mor pho log i cal char ac ter is tics of the de pos its.
This meth od olog i cal ap proach should make it pos si ble to avoid ar ti fi cially “mixed” as sem blages and to re con struct the nat u ral con di tions at the level of stages and sub-stages more pre cisely.
DISCUSSION
A “mixed” char ac ter (Kunytsia, 2007) is com monly re garded as a spe cific fea ture of the Pleis to cene land snail fau nas of the Mid dle Dniester and Ukraine. This shows the pres ence of spe - cies from dif fer ent eco log i cal groups, which to day do not co-oc - cur, in one mol lusc as sem blage. In our opin ion, such mixed as - sem blages should be in ter preted as a re sult of the di ver sity of dif fer ent stages of the Pleis to cene. It is mis lead ing to try to com - bine them while re con struct ing a sin gle land scape, since they in clude spe cies char ac ter is tic of dif fer ent nat u ral zones. The
“mixed fauna” is a sign of a mixed taphocoenosis rather than a fea ture which could dis tin guish Pleis to cene mol lusc as sem - blages from mod ern ones. The molluscs lived in dif fer ent hab i - tats but were bur ied to gether in sed i ments of the same strati - graphic unit. Their mix ing in the taphocoenosis should be con - sid ered as ev i dence of short-term in ter vals of cli mate change which could lead to changes in fau nal as sem blages. Even dur - ing the Ho lo cene, there were short warm/cold, dry/wet in ter vals (Gerasimenko, 2010b), which led to land scape changes. Such malacocoenoses could also ex ist at var i ous stages of the Pleis - to cene. The mixed as sem blages should be in ter preted as rep - re sent ing a suc ces sion fol low ing each cli mate change rather than rep re sent ing fau nas of the whole stage. This is the ap - proach we used in this pa per. To avoid the bias of “mixed fau - nas” in the study of Pleis to cene molluscs, sam pling should be done with ref er ences to well-de fined strati graphic units and must be sup ported by ac com pa ny ing meth ods (palynological, palaeopedological, ra dio car bon, etc.).
A crit i cal ap proach to palaeo eco logi cal re con struc tions of the Pleis to cene for mol lusc fau nas, which are in di ca tors not only of palaeoclimatic, but also of palaeoenvironment con di - tions, is nec es sary. Why the “palaeoenvironment”? In some cases, it is im pos si ble to in ter pret the en vi ron men tal tol er ances Tabl. 3 cont.
of cer tain spe cies of molluscs based on cli mate con di tions, as this is con tro ver sial. This is es pe cially true of Columella columella. It is a mesophile, but it is not suit able for re con struc - tions of hu mid ity lev els. As noted by Rous seau (2001), al though the Pleis to cene Pupilla as sem blages in di cate low pre cip i ta tion with in tense evap o ra tion, and the Columella as sem blage im - plies low pre cip i ta tion with al most no evap o ra tion, the re cent C.
columella lives in a tun dra-moun tain ous en vi ron ment in Scan di - na via, with an nual pre cip i ta tion of 300–473 mm. Even though this amount of pre cip i ta tion is low, the hab i tats stay damp due to the low evap o ra tion, which keeps the rocks moist and the land - scapes wa ter logged. There fore, we con sider it mis lead ing to use C. columella to in ter pret the level of pre cip i ta tion, and we re gard this spe cies only as an in di ca tor of cold con di tions.
For a more com plete in ter pre ta tion of mol lusc fau nas in dif - fer ent stages of the Pleis to cene, of their spe cies com po si tion and their spa tial dis tri bu tion, one would need to trace them along spe cific strati graphic ho ri zons and be tween dif fer ent lo - cal i ties of the Mid dle Dniester val ley, as has been done, for ex - am ple, for the site of Bullendorf in NE Aus tria (Carobene et al., 2018). How ever, due to a pres ent lack of orig i nal ma te rial, it is not yet pos si ble to so com pare the sec tions stud ied by Kunytsia with new sec tions both from the val ley of the Mid dle Dniester and with sites in the rest of Ukraine and Eu rope.
Mol lusc as sem blages var ied among dif fer ent warm and cold stages largely be cause of dif fer ent land scape and cli ma tic con di tions, with ex treme val ues of tem per a tures and hu mid ity vary ing within in di vid ual stages. Re lief also played an im por tant role, es pe cially slope ex po sure. In the study area, the Pleis to - cene de pos its are best pre served in the Dniester val ley, and mol lusc fau nas were ob tained by Kunytsia and other re search - ers only from the ter race sec tions. Mol lus can spe cies di ver sity within in di vid ual palaeo geo graphi cal stages may also have been in flu enced by sed i men tary con di tions (subaerial or subaquatic) and the dis tance from the river chan nel (low floodplain, high floodplain, low, me dium, or high ter race). This fac tor could pri mar ily af fect the pres ence or ab sence in the mol - lusc as sem blages of mois ture-sen si tive spe cies. River val leys, es pe cially deeply in cised, are also char ac ter ized by sig nif i cant land scape dif fer en ti a tion, com monly re lated to the dis tance
from the river. Even in a dry cli mate in the re gion, meso- and hygrophilous spe cies of molluscs can live on floodplains.
De spite the spe cies com po si tion di ver sity of the mol lus can fauna in dif fer ent stages of the Pleis to cene of the Mid dle Dniester, the dom i nance of some spe cies can be re cog nised.
Succinea oblonga (Drap.) was found in all stages of the Pleis to - cene. The fol low ing spe cies are also char ac ter is tic of the cold stages: Columella columella (Mart.), Vallonia tenuilabris (Al.Br.), Xerolenta obvia (Menke) [= Helicella candicans (L. Pfeif fer)). The last of these spe cies is also found in the de - pos its of al most all warm stages, ex cept for Martonosha and Zavadivka. Ap par ently, Xerolenta obvia (Menke) [= Helicella candicans (L. Pfeif fer)] was quite plas tic to the tem per a ture con - di tions dur ing the Pleis to cene pro vid ing the cli mate rel a tively arid (as a xerophilous spe cies)]. The cryophilic spe cies of cold cli ma tic cy cles have been found in the de pos its of only some warm stages: Columella columella (Mart.) only in Dofinivka time, and Vallonia tenuilabris (Al. Br.) in Pryluky and Dofinivka times. The pres ence of these and some other cryophilic spe cies and in par tic u lar, Ver tigo parcedentata (Sandb.) was ex plained by Kunytsia (2007) in terms of com plex tran si tions in time and space be tween the cold and warm stages of the Pleis to cene, which was re flected in the for ma tion of mixed, het er o ge neous fau nal as sem blages.
Dom i nant spe cies of the cold stages in clude Ver tigo parcedentata (Sandb.), Pupilla sterri (Voith), Helicopsis striata (Müll.) and Pseudotrichia rubiginosa (Al. Schm.), which are char ac ter is tic of most, though not all, cold stages. The fol low ing spe cies are also quite com mon: Pupilla triplicata (Studer), Chondrula tridens (Müll.) and Euconulus fulvus (Müll.).
Helicopsis instabilis (Rossmässler) and Trichia hispida (L.), char ac ter is tic of the last cold stages (Uday, Bug and Prychornomoria). The pres ence of the lat ter two spe cies in ear - lier cold climatolithes seems not to have been noted (Kunytsia, 1965, 1966, 1971, 1974, 1984, 2007; Motuz, 1977, 1982, 1987;
Melnychuk, 1984, 2004; Dmytruk, 2004; Ivchenko, 2010;
Bogucki et al., 2011, 2012; Alexandrowicz et al., 2014).
Groups of molluscs of warm Pleis to cene climatolithes of the Mid dle Dniester dif fer sig nif i cantly in spe cies com po si tion, so fewer dom i nant spe cies are found here. Succinea oblonga (Drap.), noted above, is found at all stages. Taxa such as Vallonia pulchella (Müll.) and Chondrula tridens (Müll.) have not been iden ti fied only for the Zavadika and Dofinivka stages.
Quite com mon are Pupilla sterri (Voith), Clausilia dubia (Drap.), Xerolenta obvia (Menke) (= Helicella candicans (L. Pfeif fer), Helicopsis striata (Müll.), Trichia hispida (L.), Arianta arbustorum (L.), but these taxa are not char ac ter is tic of all stages.
The fol low ing taxa are mod er ately rare in the Up per and Mid dle Pleis to cene de pos its of the Mid dle Dniester Val ley:
Vallonia excentrica Sterki, Ver tigo arctica (Wall.), V. pseudosubstriata Ložek, Cochlodina laminata (Montagu), Clausilia bidentata (Ström), Laciniaria plicata (Drap.), Punctum pygmaeum (Drap.), Vitrea crystallina (Müll.), Aegopinella nitens (Michaud), Nesovitrea pet ro nella (L. Pfeif fer), Euomphalia strigella (Drap.).
In gen eral, biostratigraphic anal y sis of the Pleis to cene molluscs of the Mid dle Dniester showed that the mol lusc as - sem blages of the cold stages are more di verse than in the warm stages (Fig. 2). The high est spe cies di ver sity was es tab lished for the Bug climatolith. It is also high for the Sula and Prychornomoria stages. Among the warm stages, the high est spe cies di ver sity is char ac ter is tic for the last three: Pryluky, Vytachiv and Dofinivka. For both cold and warm stages, high spe cies di ver sity is ob served at the end of the Pleis to cene (Pryluky–Prychornomoria stages).
Fig. 2. Num ber of ter res trial mol lusc spe cies in the Mid dle and Late Pleis to cene stages in the Mid dle Dniester area Hor i zon tally: num ber of spe cies; ver ti cally: Pleis to cene stages (see the in di ces in Ta ble 1); * – stages not re corded in the area
stud ied
CONCLUSIONS
Anal y sis of mol lusc fau nas from dif fer ent stages of the Pleis to cene in the Mid dle Dniester area shows that the spe cies and eco log i cal com po si tion of the as sem blages vary from stage to stage. The dif fer ences in the as sem blages’ spe cies com po si - tion re flect the spa tial and tem po ral changes of the palaeolandscapes and as well as global and lo cal cli mate changes. Ac cord ingly, each stage of the Pleis to cene has its own in di vid ual tax o nomic and eco log i cal set of spe cies.
The mol lusc as sem blages of those stages in which the en vi - ron men tal con di tions were par tic u larly change able are more var ied. The changes in volved two fac tors – air tem per a ture (warm/cold phases) and hu mid ity (wet/dry). Such changes, in the first place, led to land scape changes and, con se quently, af - fected the spe cies com po si tion and ecol ogy of the mol lusc fau - nas, which now makes it pos si ble to in ter pret those changes.
Ana lys ing the fauna of the cold and warm stages of the Pleis to cene, we found that the lat ter were more di verse. While
dur ing the cold stages there were many spe cies in com mon for all or most stages, the fau nas of the warm stages were more var ied. Such a fea ture can play an im por tant role in Qua ter nary biostratigraphy.
The biodiversity and spe cific fea tures of the Mid dle Dniester mol lusc as sem blages fa cil i tate un der stand ing of the evo lu tion of en vi ron men tal con di tions dur ing the Pleis to cene and make it pos si ble to iden tify in di vid ual en vi ron men tal events re flected by re sponses among the mol lusc com mu ni ties. These re sults can serve as a pow er ful in stru ment in re gional palaeogeographic anal y sis.
Ac knowl edge ments. This work was partly sup ported by INQUA pro ject 1606P “Ground squir rels on the march: ex pan - sion and speciation in the Qua ter nary of the Circum-Pontic area and sur round ings”. We thank two anon y mous re view ers for their con struc tive re view of the manu script. We ex tend our grat - i tude to B. Pokryszko for proof read ing the text.
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